Several taxa of the Australian Proteaceae have invaded South African fynbos and require costly management programmes to bring under control. Banksia spp. have been introduced only recently to fynbos regions. The invasive potential of Banksia ericifolia (Proteaceae) was investigated by comparing its recruitment potential with that of an indigenous proteaceous shrub, Leucadendron laureolum. Both species are overstorey shrubs that are killed by fire and rely on canopy-stored seeds (serotiny) for recruitment. Eight year old B. ericifolia shrubs produced an average of 16 500 seeds per plant, which is thirty times more than the average of 570 seeds produced by 10 year old L. laureolum shrubs. The seed bank of B. ericifolia was not only larger than that of L. laureolum (1098 vs 525 viable seeds m−2 projected canopy cover), but also considerably larger than that described for the species in its native environment (200–330 seeds m−2 in a 9 year old stand north of Sydney). Leucadendron laureoleum released most of its seed a few days after the cones were burnt, whereas seed release in B. ericifolia was spread over 12 weeks. The seeds of B. ericifolia had lower wingloading and fall rates than L. laureolum and were dispersed over greater distances. The relative seedling growth rates of the two species were very similar (0.03 g per day), but below-ground biomass was greater and proteoid roots were more developed in B. ericifolia seedlings than in L. laureolum after 100 days. Four year old B. ericifolia plants growing in the field had attained over twice the height of indigenous pro-teoids and accumulated up to 10 times the fresh biomass of L. xanthoconus, a species which is ecologically similar to L. laureolum. The Bioclimatic Prediction System (BIOCLIM) was used to create a bioclimatic profile of B. ericifolia and identify climatically suitable areas in the Cape Province. Results show that its potential distribution covers most fynbos areas in the southwestern Cape. It is concluded that B. ericifolia has the potential to be highly invasive in fynbos.

The invasive potential of Australian banksias in South African fynbos: A comparison of the reproductive potential of Banksia ericifolia and Leucadendron laureolum

Proteaceae are most diverse in southern Africa and Australia, especially in the south-western portions of these regions. Most genera have some species in flower at all times of the year, although generally there is a preponderance of species that flower between late winter and early summer. The one genus that is an exception to this generalization is Banksia, which either has approximately the same percentage of species in flower at various times of the year (southwestern Australia) or peaks in autumn (southeastern Australia). Within particular communities, opportunities for hybridization among congeneric species are minimized by staggered flowering times, different pollen vectors and/or various incompatibility mechanisms. Birds, mammals and arthropods have been identified as visitors to the inflorescences of many Proteaceae. The most common avian visitors to the majority of genera in Australia are honeyeaters, although lorikeets, silvereyes and approximately 40 other species sometimes may be important. Sugarbirds and sunbirds are seen most frequently at inflorescences of Protea, Leucospermum and Mimetes in southern Africa, although they rarely visit other genera. In most cases, avian visitors forage in a manner that permits the acquisition and transfer of pollen. Limited evidence supports the hypothesis that birds are selective in their choice of inflorescences, responding to morphological and/or colour changes and usually visiting those inflorescences that offer the greatest nectar rewards. Arthropods may be equally selective, although it is possible that only the larger moths, bees and beetles are important pollinators, even for those plant species that rely entirely on arthropods for pollen transfer. Mammals are pollen vectors for some Proteaceae, especially those that have geoflorous and/or cryptic inflorescences. In Australia, small marsupials may be the most important mammalian pollinators, although rodents fill this niche in at least some southern African habitats. All but two genera of Proteaceae are hermaphroditic and protandrous, the exceptions being the dioecious southern African genera Aulax and Leucadendron. For hermaphroditic species, the timing of visits by animals to inflorescences is such that they not only acquire pollen from freshly opened flowers but also brush against pollen presenters and stigmas of others that have lost self-pollen and become receptive. Birds and insects (and probably mammals) generally forage in such a way as to facilitate both outcrossing and selfing. Some species are self-compatible, although many require outcrossing if viable seed is to be formed. Regardless of which animals are the major pollen vectors, fruit set is low relative to the number of flowers available, especially in Australian habitats. Functional andromonoecy of the majority of flowers is advanced as the major cause of poor fruit set. The pollination biology and breeding systems of Australian and southern African Proteaceae resemble one another in many ways, partly because of their common ancestry, but also due to convergence. Divergence is less obvious, apart from the dichotomy between dioecious and hermaphroditic genera, and differences in the levels of seed set for Australian and African species. Future studies should concentrate on identifying the most important pollinators for various Proteaceae, the manner in which their visits are integrated with floral development and factors responsible for limiting fruit set.

Pollination biology of the Proteaceae in Australia and southern Africa

gan 200 years ago, when Europeans settled on the continent. With that colonization, natural habitats were cleared and degraded, and a wide variety of domesticated animals and plants were introduced (Saunders et al. 1990). One of the animals introduced was the honeybee, Apis mellifera. This social bee arrived approximately 150 years ago (Doull 1973), and now feral colonies and colonies managed by beekeepers are spread over most of Australia.

Honeybees in the Australian EnvironmentDoes Apis mellifera disrupt or benefit the native biota

The number of fruits produced by many hermaphroditic plants is usually far fewer than the number of flowers available for fertilization. There are various possible explanations for the low fruit:flower ratio, some proximate and others ultimate. Recent studies, especially in northern hemisphere systems, have used field experiments to test some of them, but there are potential difficulties with the methodology of some experiments and with the testing of ultimate hypotheses. It is important to examine the possible explanations for low fruit: flower ratios with a range of different systems. This article reviews studies on Australian species of woody, perennial shrubs in the family Proteaceae; this evolutionarily distinct group of plants and pollinators has several unusual and interesting characteristics, and provides a valuable addition to the better-known northern hemisphere studies.

Factors controlling fruit set in hermaphroditic plants: Studies with the Australian proteaceae

European honeybees (Apis mellifera) were less efficient pollinators ofGrevillea barklyana than nectar-feeding birds. Nectar-collecting honeybees did not contact reproductive parts of flowers. Pollen-collecting honeybees preferentially visited malestage flowers but rarely visited female-stage flowers. Fruit set on caged inflorescences that allowed access to honeybees but excluded birds was reduced by more than 50% compared to inflorescences that were visited by both types of visitors. Further, fruit set on caged inflorescences was less than on bagged inflorescences that excluded both birds and honeybees, indicating that pollen removal by bees decreased opportunities for delayed autonomous selfing in the absence of birds. Although fruit set was not pollen-limited at the study site, pollen removal by honeybees would decrease fruit set in small populations where birds are scarce. In addition, pollen removal by honeybees would reduce opportunities for outcrossing and reproductive success through male function. Although honeybees have been in Australia for insufficient time to have exerted selection on floral traits, evolutionary shifts in response to these animals are likely to occur in the future.

Pollination disruption by European honeybees in the Australian bird-pollinated shrub Grevillea barklyana ( Proteaceae )

In coastal heath, 12 km north of Sydney, Banksia ericifolia set fruit after cross-pollination but not after controlled self-pollination. Animals removed nectar and pollen from inflorescences during the day but not overnight. Introduced honeybees (Apis mellifera) and native birds (Meliphagidae, Zosteropidae) visited flowers frequently during the day, carried pollen on their body surfaces and were likely to transfer pollen between plants. Native bees, nocturnal moths and ants also visited flowers but were too infrequent, did not move between plants or foraged inappropriately to be important pollinators. No mammals were seen visiting flowers and the small mammals we caught carried no pollen. Exclusion experiments confirmed that pollination occurred during the day and not at night, but the experiments were insufficient to determine the relative importance of honeybees versus birds as pollinators. Only about 3% of the flowers of Banksia ericifolia developed follicles under natural conditions. Follicle production in this species was limited by resources and not by pollinators, since multiple cross- pollinations of flowers did not increase follicle production above the natural rate.

Pollination of Banksia ericifolia Smith: Birds, Mammals and Insects as Pollen Vectors

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Pollination of Banksia ericifolia Smith: Birds, Mammals and Insects as Pollen Vectors